In the literature, litter size in red squirrels Sciurus vulgaris ranges between
1-6 offspring. Here we report a record litter size, in Europe, of 7 young (6 males, 1 female),
marked in the nest when 10-14 days old, in a high-elevation mountain pine habitat, Lombardy
Alps (Northern Italy). We discuss implications of extreme winter conditions for lifehistory
strategies and possible trade-offs between high reproductive investment and short
life-spa

Mari, V.

Martini, S.

Martinoli, A.

Molinari, A.

Romeo, C.

Tosi, G.

Wauters, L.A.

In the literature, litter size in red squirrels Sciurus vulgaris ranges between
1-6 offspring. Here we report a record litter size, in Europe, of 7 young (6 males, 1 female),
marked in the nest when 10-14 days old, in a high-elevation mountain pine habitat, Lombardy
Alps (Northern Italy). We discuss implications of extreme winter conditions for lifehistory
strategies and possible trade-offs between high reproductive investment and short
life-spa

MARTINOLI, ADRIANO

TOSI, GUIDO

WAUTERS, LUCAS ARMAND

Archivio istituzionale della ricerca - Università dell'Insubria

Hystrix It. J. Mamm. (n.s.) 19 (1) (2008): 61-65  RECORD LITTER SIZE IN THE EURASIAN RED SQUIRREL (SCIURUS VULGARIS)  VERONICA MARI, STEFANIA MARTINI, CLAUDIA ROMEO, AMBROGIO MOLINARI, ADRIANO MARTINOLI, GUIDO TOSI, LUCAS A. WAUTERS*  Dipartimento Ambiente, Salute, Sicurezza, Università degli Studi dell’Insubria, Varese, Via J.H. Dunant 3, I-21100 Varese, Italy *Corresponding author, e-mail: l.wauters@uninsubria.it  Received 2 October 2007; accepted 15 January 2008  ABSTRACT - In the literature, litter size in red squirrels Sciurus vulgaris ranges between 1-6 offspring. Here we report a record litter size, in Europe, of 7 young (6 males, 1 female), marked in the nest when 10-14 days old, in a high-elevation mountain pine habitat, Lom-bardy Alps (Northern Italy). We discuss implications of extreme winter conditions for life-history strategies and possible trade-offs between high reproductive investment and short life-span.  Key words: Sciurus vulgaris, litter size, life-history tactics, marginal habitat, mountain pine forest  RIASSUNTO - Dimensione eccezionale della cucciolata di scoiattolo comune (Sciurus vulgaris). Dai dati di letteratura, il numero di esemplari per cucciolata nello scoiattolo co-mune Sciurus vulgaris varia generalmente da 1 a 6. Nel presente lavoro viene riportata per la prima volta in Europa la segnalazione di una cucciolata di 7 esemplari (6 maschi e 1 femmina), che sono stati marcati nel nido all’età di 10-14 giorni. Il nido è stato individuato in un habitat forestale dominato da pino mugo nel Parco Nazionale dello Stelvio, in Lom-bardia. Vengono prese in esame le implicazioni che le condizioni estreme, nel periodo in-vernale, possono avere sulle strategie riproduttive dello scoiattolo comune.  Parole chiave: Sciurus vulgaris, dimensione cucciolata, strategia riproduttiva, habitat mar-ginale, foresta montana di conifere  INTRODUCTION  In food-limited habitats, the energy transferred by a lactating female to her offspring is limited by her ability to lo-cate and ingest sufficient amounts of food prior to (capital breeders) or dur-ing (income breeders) lactation (Reznick, 1992; Weiner, 1992; Hum-phries and Boutin, 2000). Under con-straints of food availability, physiolo-gical energetic ceilings and life-history trade-offs (e.g. smaller litters have higher individual survival than larger ones), a breeding female should pro-duce the number and size of offspring, each breeding season, that maximises its lifetime reproductive success (Wil-liams, 1966; Clutton-Brock, 1988; Humphries and Boutin, 2000). 61Mari et al.  Red squirrels (Sciurus vulgaris) can be considered income breeders since sea-sonal reproductive success depends on the amount of (high-quality) food to which a breeding female has access (Wauters and Dhondt, 1989, 1995; Wauters and Lens, 1995). Variation in lifetime reproductive success among females is mainly determined by varia-tion in breeding lifespan (thus age of first reproduction, lifespan), but also by differences in litter size (measured in most cases around weaning-age, 8-12 weeks, Wauters et al., 1993; Wauters and Dhondt, 1995; Wauters and Lens, 1995), and the probability of juve-niles/subadults to settle success-sfully (Wauters and Dhondt, 1993, 1995; Wauters et al., 1993). The latter de-pends on body mass differences, with heavier young being more likely to set-tle than those of lower body mass, which is partly affected by the mother’s mass at the onset of breeding (Wauters and Dhondt, 1993; Wauters et al., 1993). Average litter sizes, at weaning, also differ between years in relation to seed-crop size of the major feeding trees (Wauters and Lens, 1995; Boutin et al., 2006).  Here we report an exceptionally large litter size, never recorded in any field study on free-ranging red squirrels in Europe.  STUDY AREA  Squirrels were studied in the central Italian Alps in a dwarf mountain pine (Pinus mugo) woodland with trees with both pros-trate (3%) and arboreal habit (96%), nearby the artificial Cancano lakes (46°32’ N, 10°16’ E) in the Stelvio National Park. The study area covers 54 ha at a mean elevation of 1950 m. a.s.l. The area is almost entirely composed of a dense mountain pine forest (average tree density 3308 trees/ha) with few dead trees (1%). Average diameter at breast height is 14 cm. The undergrowth is composed of heather (Erica carnea), juniper (Juniperus com-munis), blueberry (Vaccin-ium myrtillus) and lingonberry (Vaccinium vitis - idea). The mountain pine habitat is at the upper-limit of the squirrel’s distribution and is character-rised by long and cold win-ters (e.g. winter November 2005-April 2006: mean monthly minimum temperature from -11.7°C in January to -2.7°C in April; mean monthly maximum temperature from -2.6°C in December to 6.7°C in April), with perma-nent, but patchy snow cover between De-cember and mid to late April, with marked annual variation in snow depth.  METHODS  Squirrels were live-trapped in three sepa-rate sessions per year, starting March-April, June-July and September-October. A trapping session involved the use of 20 to 30 ground-placed Tomahawk traps (models 201 and 202, Tomahawk Live Trap Co., Wisconsin, USA). Traps were placed on a grid, with distances of 100-150 m between traps and average trap density of 0.6-0.7 traps ha-1. Traps were pre-baited with sun-flower seeds and hazelnuts four to six times over a 30-day period, and then baited and set for eight to 12 days, until no new, un-marked squirrels were trapped for at least two consecutive days. Traps were partly covered by dark plastic to give shelter from rain or cold, and checked two-three times per day. Each trapped squirrel was flushed into a light cotton handling bag with a zip-per (Wauters et al., 2007), or a wire-mesh ‘handling cone’ to minimise stress during handling, and individually marked using numbered metal ear-tags (type 1003 S, 10 by 2 mm, National Band and Tag Co, Newport, Kentucky, USA). It was weighed to the nearest 5 g using a Pesola spring-balance (Pesola AG, Baar, Switzerland), 62Record litter size in red squirrel  and the length of the right hind foot (with-out nail) was measured (0.5 mm) with a thin ruler (Wauters et al., 2007). Sex, age, and reproductive condition were recorded following Wauters and Dhondt (1995). At first capture a skin sample was taken for DNA analysis using a biopsy punch of 3 mm diameter (Stiefel laboratories Milano, Italy). All squirrels first caught as juveniles (6 - 15 weeks old, body mass below 250g, Wauters et al., 2007) were defined as lo-cally born offspring (see also Wauters and Dhondt, 1995). Moreover, by radio-tracking adult females in 2007 (Wauters and Dhondt, 1995; Wauters and Lens, 1995; Wauters et al., 2005), we were able to locate the nursing nest of two lactating females and tag their offspring (Wauters et al., 1993; Wauters and Dhondt, 1995). These juveniles were taken by hand from the nest when the female was absent (as revealed by radio-transmitter signal), han-dled, sexed and marked on the ground, and delicately put back in the nest. To minimise stress and handling time we did not weigh them. The total operation of climbing the tree, taking and handling the young, and returning them to the nest took less than 20 minutes. The next day, we checked whether the mother still nursed the young in the same nest or whether she had moved them to a new nest. None of the litters was abandon-ned by the female (Wauters et al., 1993; Wauters and Dhondt 1995; Wauters and Lens 1995; Humphries and Boutin 2000; Boutin et al., 2006).  RESULTS AND DISCUSSION  Female F2340 was radio-tagged on 9 March 2007, weighing 295g, when in pre-oestrus conditions. She was recap-tured pregnant on 24 April (body mass 330g) when palpation of the abdomen revealed the presence of three embryos. The nursing nest was found on 16 May and the three young (1 male, 2 females) were marked on 31 May 2007. Young were still blind, had already a dense coat and their lower incisors just started to erupt. Based on these characteristics their age was estimated at 20-25 days (Eibl-Eibesfeldt, 1951; Tittensor, 1977). On 26 June, when offspring was about 7 weeks old, the female was re-captured and still lactating, indicating that she successfully raised her litter. Body mass was 325g. On 22 June 2007, F2167 was marked and radio-collared. She was pregnant and weighed 350g. The nursing nest was found on 27 July and seven young (6 males, 1 female) were marked. Off-spring was blind and had only a downy fur cover emerging, typical for 10-14 day-old squirrels (Eibl-Eibe-sfeldt, 1951; Tittensor, 1977). A litter of seven was the largest one ever recorded for red squirrels (but see Ognev, 1940). In the UK, average litter size between birth and first weeks of life was 3.2 young (N = 81, Tittensor, 1977), and reviews reported litter sizes ranging from 2-5 (Lange et al., 1994) and 1-4, max. 6 young (Tittensor, 1977; Lurz et al., 2005). We only found a single cita-tion from the (translated) Russian lit-erature on “10 squirrels found in a nest” (in Ognev, 1940), but it is not clear whether this refers to a single lit-ter. Detailed capture-mark-recapture studies in Western Europe reported maximum litter size (between birth and 1-week old) of four and average litter size of 3.0 ± 0.9 (N = 9, Wauters and Lens, 1995). Finally, even in a pine wood with supplementary feeding, av-erage litter size was not higher than in natural populations (mean ± S.D. = 3.17 ± 0.47 and 2.40 ± 0.31 young in 63Mari et al.  two consecutive years, Shuttleworth, 1996 cited in Lurz et al., 2005). Off-spring losses during lactation can be high, and even complete litters can be lost due to predation or when body condition of the mother is reduced (loss of body mass between birth and wean-ing, Wauters and Dhondt, 1989, 1995; Wauters and Lens, 1995; Lurz et al., 2005). In the case of the litter of seven, two juveniles were observed at less than 50m from the nest tree on August 31st and September 1st (ca. 6-7 weeks old), foraging on pine cones. One of them, a male, was recaptured at the end of October (ca. 3.5 months old), when also a male of the litter of three was trapped (nearly 6 months old). Finally, trapping revealed that the other two resident adult females (F2359, body mass 295g, and F2429, body mass 395g) also produced a litter (caught in lactation on August 31st and September 2nd , respectively) and three juveniles were caught and marked on 1 and 2 September 2007 when about 10-12 weeks old (200 – 230g). One of the females probably had two litters in 2007, since we also captured a small juvenile (100g, ca. 6-7 weeks old) on 25 June 2007. This will be tested using parent assignment tests on DNA sam-ples. Overall offspring sex-ratio (10 males, 4 females) was not different from even (χ21 = 2.57; P = 0.11). Body mass differed strongly among breeding females (range 295 – 395g), but two of them had a high lactation mass for fe-male squirrels in populations through-out the Alps (Wauters et al., 2007). How can we explain this extremely high reproductive rate (14 offspring from 4 adult females = 3.5 juv./reproducing female)? We suggest that in this extreme habitat, with strong annual fluctuations in seed-crop size and thus food availability and long and cold winters with deep snow-cover, winter mortality can be very high. This was confirmed by our preliminary data (winter survival September 2006 - March 2007 = 0.25). Hence, the prob-ability that squirrels only have one or two years to reproduce is high and fe-males should adopt a trade-off between high reproductive investments when food abundance is high, against re-duced survival (Williams, 1966; Clut-ton-Brock, 1988). We therefore predict that in mountain pine forests, red squir-rels are selected to reproduce as year-lings and to invest heavily in reproduc-tion in the first year after birth they have access to a rich seed-crop. To test this hypothesis, we will continue to monitor annual seed-crop production, squirrel numbers, reproductive success of all resident females and offspring survival/recruitment over the following years.  ACKNOWLEDGEMENTS  We thank AEM company for logistic support, and Matteo Wauters for company and tree climbing. Construc-tive criticism by two referees helped to improve the manuscript. This study is part of the ASPER (Alpine Squirrel Population Ecology Research) project, funded by the province of Sondrio (Servizio Agricoltura, Caccia e Pesca, Settore Risorse Ambientali) and the Stelvio National Park to Istituto Oikos NGO, Milan. Additional financial support was given by MIUR (Ministero dell’Istruzione, dell’Università della ricerca, project COFIN 2003, number 64Record litter size in red squirrel  2003053710-006) to Insubria Universi-ty of Varese. This is paper12 of the ASPER project.  REFERENCES  Boutin S., Wauters L.A., McAdam A.G., Humphries M.M., Tosi G. and Dhondt A.A. 2006. Anticipatory reproduction and population growth in seed preda-tors. Science, 314: 1928-1930. Clutton-Brock T.H. 1988. Reproductive success, University of Chicago Press, Chicago. Eibl-Eibesfeldt I. 1951. Beobachtungen zur Fortplanzungsbiologie und Jugendent-wicklung des Eichhörnchens (Sciurus vulgaris L.). Z. Tierpsychol., 8: 370-400. Humphries M.M. and Boutin S. 2000. The determinants of optimal litter size in free-ranging red squirrels. Ecology, 81: 2867-2877. Lange R., Twisk P., Van Winden A. and van Diepenbeek A. 1994. Zoogdieren van West-Europa, KNNV-Uitgeverij, Utrecht, pp. 88-92. (in Dutch) Lurz P.W.W., Gurnell J., and Magris L. 2005. Sciurus vulgaris. Mammalian Species, American Society of Mamma-logists, 769: 1-10 Ognev S. I. 1940. Mammals of the U.S.S.R. and adjacent countries. Mam-mals of eastern Europe and northern Asia. Translated from Russian, Israel programme for scientific translations, Jerusalem, 1966, Vol IV: 284-377.  Reznick D. 1992. Measuring the cost of reproduction. Trends Ecol. Evol., 7: 42-45. Tittensor A.M. 1977. Red Squirrel. In: Corbet G.B., Southern H.N (eds.), The Handbook of British Mammals, 2nd edition. Blackwell Scientific Publica-tions, Oxford, 153-164. Wauters L.A. and Dhondt A.A. 1989. Body weight, longevity and reproductive success in red squirrels (Sciurus vulga-ris). J. Anim. Ecol., 58: 637-651. Wauters L.A., and Dhondt A.A. 1993. Im-migration patterns and success in red squirrels. Behav. Ecol. Sociobiol., 33: 159-167. Wauters L.A. and Dhondt A.A. 1995. Components of lifetime reproductive success of female Eurasian red squir-rels. Oikos, 72: 402-410. Wauters L.A. and Lens L. 1995. Effects of food availability and density on red squirrel (Sciurus vulgaris) reprodu-ction. Ecology, 76: 2460-2469. Wauters L.A., Bertolino S., Adamo M., Van Dongen S. and Tosi G. 2005. Food shortage disrupts social organiza-tion: the case of red squirrels in conifer forests. Evol. Ecol., 19: 375-404. Wauters L.A., Bijnens L. and Dhondt A.A. 1993. Body mass at weaning and juve-nile recruitment in the red squirrel. J. Anim. Ecol., 62: 280-286. Wauters L.A., Vermeulen M., van Dongen S., Bertolino S., Molinari A., Tosi G. and Matthysen E. 2007. Effects of spa-tio-temporal variation in food supply on red squirrel Sciurus vulgaris body size and body mass and its conse-quences for some fitness com-ponents. Ecography, 30: 51-65. Weiner J. 1992. Physiological limits to sus-tainable energy budgets in birds and mammals: ecological implications. Trends Ecol. Evol., 7: 384-388. Williams G.C. 1966. Natural selection, the costs of reproduction, and a refinement of Lack’s principle. Am. Nat., 100: 687-690.    65

Record litter size in the eurasian red squirrel (Sciurus vulgaris)

In the literature, litter size in red squirrels Sciurus vulgaris ranges between 1-6 offspring. Here we report a record litter size, in Europe, of 7 young (6 males, 1 female), marked in the nest when 10-14 days old, in a high-elevation mountain pine habitat, Lombardy Alps (Northern Italy). We discuss implications of extreme winter conditions for lifehistory strategies and possible trade-offs between high reproductive investment and short life-span

V. Mari

S. Martini

C. Romeo

A. Molinari

A. Martinoli

G. Tosi

L.A. Wauters

AIR Universita degli studi di Milano

Hystrix It. J. Mamm. (n.s.) 19 (1) (2008): 61-65 
 
RECORD LITTER SIZE IN THE EURASIAN RED SQUIRREL 
(SCIURUS VULGARIS) 
 
VERONICA MARI, STEFANIA MARTINI, CLAUDIA ROMEO, AMBROGIO 
MOLINARI, ADRIANO MARTINOLI, GUIDO TOSI, 
LUCAS A. WAUTERS* 
 
Dipartimento Ambiente, Salute, Sicurezza, Università degli Studi dell’Insubria, Varese, Via 
J.H. Dunant 3, I-21100 Varese, Italy 
*Corresponding author, e-mail: l.wauters@uninsubria.it 
 
Received 2 October 2007; accepted 15 January 2008 
 
ABSTRACT - In the literature, litter size in red squirrels Sciurus vulgaris ranges between 
1-6 offspring. Here we report a record litter size, in Europe, of 7 young (6 males, 1 female), 
marked in the nest when 10-14 days old, in a high-elevation mountain pine habitat, Lom-
bardy Alps (Northern Italy). We discuss implications of extreme winter conditions for life-
history strategies and possible trade-offs between high reproductive investment and short 
life-span. 
 
Key words: Sciurus vulgaris, litter size, life-history tactics, marginal habitat, mountain pine 
forest 
 
RIASSUNTO - Dimensione eccezionale della cucciolata di scoiattolo comune (Sciurus 
vulgaris). Dai dati di letteratura, il numero di esemplari per cucciolata nello scoiattolo co-
mune Sciurus vulgaris varia generalmente da 1 a 6. Nel presente lavoro viene riportata per 
la prima volta in Europa la segnalazione di una cucciolata di 7 esemplari (6 maschi e 1 
femmina), che sono stati marcati nel nido all’età di 10-14 giorni. Il nido è stato individuato 
in un habitat forestale dominato da pino mugo nel Parco Nazionale dello Stelvio, in Lom-
bardia. Vengono prese in esame le implicazioni che le condizioni estreme, nel periodo in-
vernale, possono avere sulle strategie riproduttive dello scoiattolo comune. 
 
Parole chiave: Sciurus vulgaris, dimensione cucciolata, strategia riproduttiva, habitat mar-
ginale, foresta montana di conifere 
 
INTRODUCTION 
 
In food-limited habitats, the energy 
transferred by a lactating female to her 
offspring is limited by her ability to lo-
cate and ingest sufficient amounts of 
food prior to (capital breeders) or dur-
ing (income breeders) lactation 
(Reznick, 1992; Weiner, 1992; Hum-
phries and Boutin, 2000). Under con-
straints of food availability, physiolo-
gical energetic ceilings and life-history 
trade-offs (e.g. smaller litters have 
higher individual survival than larger 
ones), a breeding female should pro-
duce the number and size of offspring, 
each breeding season, that maximises 
its lifetime reproductive success (Wil-
liams, 1966; Clutton-Brock, 1988; 
Humphries and Boutin, 2000). 
61
Mari et al. 
 
Red squirrels (Sciurus vulgaris) can be 
considered income breeders since sea-
sonal reproductive success depends on 
the amount of (high-quality) food to 
which a breeding female has access 
(Wauters and Dhondt, 1989, 1995; 
Wauters and Lens, 1995). Variation in 
lifetime reproductive success among 
females is mainly determined by varia-
tion in breeding lifespan (thus age of 
first reproduction, lifespan), but also by 
differences in litter size (measured in 
most cases around weaning-age, 8-12 
weeks, Wauters et al., 1993; Wauters 
and Dhondt, 1995; Wauters and Lens, 
1995), and the probability of juve-
niles/subadults to settle success-sfully 
(Wauters and Dhondt, 1993, 1995; 
Wauters et al., 1993). The latter de-
pends on body mass differences, with 
heavier young being more likely to set-
tle than those of lower body mass, 
which is partly affected by the mother’s 
mass at the onset of breeding (Wauters 
and Dhondt, 1993; Wauters et al., 
1993). Average litter sizes, at weaning, 
also differ between years in relation to 
seed-crop size of the major feeding 
trees (Wauters and Lens, 1995; Boutin 
et al., 2006).  
Here we report an exceptionally large 
litter size, never recorded in any field 
study on free-ranging red squirrels in 
Europe. 
 
STUDY AREA 
 
Squirrels were studied in the central Italian 
Alps in a dwarf mountain pine (Pinus 
mugo) woodland with trees with both pros-
trate (3%) and arboreal habit (96%), nearby 
the artificial Cancano lakes (46°32’ N, 
10°16’ E) in the Stelvio National Park. The 
study area covers 54 ha at a mean elevation 
of 1950 m. a.s.l. The area is almost entirely 
composed of a dense mountain pine forest 
(average tree density 3308 trees/ha) with 
few dead trees (1%). Average diameter at 
breast height is 14 cm. The undergrowth is 
composed of heather (Erica carnea), juniper 
(Juniperus com-munis), blueberry (Vaccin-
ium myrtillus) and lingonberry (Vaccinium 
vitis - idea). The mountain pine habitat is at 
the upper-limit of the squirrel’s distribution 
and is character-rised by long and cold win-
ters (e.g. winter November 2005-April 2006: 
mean monthly minimum temperature from -
11.7°C in January to -2.7°C in April; mean 
monthly maximum temperature from -2.6°C 
in December to 6.7°C in April), with perma-
nent, but patchy snow cover between De-
cember and mid to late April, with marked 
annual variation in snow depth. 
 
METHODS 
 
Squirrels were live-trapped in three sepa-
rate sessions per year, starting March-
April, June-July and September-October. A 
trapping session involved the use of 20 to 
30 ground-placed Tomahawk traps (models 
201 and 202, Tomahawk Live Trap Co., 
Wisconsin, USA). Traps were placed on a 
grid, with distances of 100-150 m between 
traps and average trap density of 0.6-0.7 
traps ha-1. Traps were pre-baited with sun-
flower seeds and hazelnuts four to six times 
over a 30-day period, and then baited and 
set for eight to 12 days, until no new, un-
marked squirrels were trapped for at least 
two consecutive days. Traps were partly 
covered by dark plastic to give shelter from 
rain or cold, and checked two-three times 
per day. Each trapped squirrel was flushed 
into a light cotton handling bag with a zip-
per (Wauters et al., 2007), or a wire-mesh 
‘handling cone’ to minimise stress during 
handling, and individually marked using 
numbered metal ear-tags (type 1003 S, 10 
by 2 mm, National Band and Tag Co, 
Newport, Kentucky, USA). It was weighed 
to the nearest 5 g using a Pesola spring-
balance (Pesola AG, Baar, Switzerland), 
62
Record litter size in red squirrel 
 
and the length of the right hind foot (with-
out nail) was measured (0.5 mm) with a 
thin ruler (Wauters et al., 2007). Sex, age, 
and reproductive condition were recorded 
following Wauters and Dhondt (1995). At 
first capture a skin sample was taken for 
DNA analysis using a biopsy punch of 3 
mm diameter (Stiefel laboratories Milano, 
Italy). All squirrels first caught as juveniles 
(6 - 15 weeks old, body mass below 250g, 
Wauters et al., 2007) were defined as lo-
cally born offspring (see also Wauters and 
Dhondt, 1995). Moreover, by radio-
tracking adult females in 2007 (Wauters 
and Dhondt, 1995; Wauters and Lens, 
1995; Wauters et al., 2005), we were able 
to locate the nursing nest of two lactating 
females and tag their offspring (Wauters et 
al., 1993; Wauters and Dhondt, 1995). 
These juveniles were taken by hand from 
the nest when the female was absent (as 
revealed by radio-transmitter signal), han-
dled, sexed and marked on the ground, and 
delicately put back in the nest. To minimise 
stress and handling time we did not weigh 
them. The total operation of climbing the 
tree, taking and handling the young, and 
returning them to the nest took less than 20 
minutes. 
The next day, we checked whether the 
mother still nursed the young in the same 
nest or whether she had moved them to a 
new nest. None of the litters was abandon-
ned by the female (Wauters et al., 1993; 
Wauters and Dhondt 1995; Wauters and 
Lens 1995; Humphries and Boutin 2000; 
Boutin et al., 2006). 
 
RESULTS AND DISCUSSION 
 
Female F2340 was radio-tagged on 9 
March 2007, weighing 295g, when in 
pre-oestrus conditions. She was recap-
tured pregnant on 24 April (body mass 
330g) when palpation of the abdomen 
revealed the presence of three embryos. 
The nursing nest was found on 16 May 
and the three young (1 male, 2 females) 
were marked on 31 May 2007. Young 
were still blind, had already a dense 
coat and their lower incisors just started 
to erupt. Based on these characteristics 
their age was estimated at 20-25 days 
(Eibl-Eibesfeldt, 1951; Tittensor, 
1977). On 26 June, when offspring was 
about 7 weeks old, the female was re-
captured and still lactating, indicating 
that she successfully raised her litter. 
Body mass was 325g. 
On 22 June 2007, F2167 was marked 
and radio-collared. She was pregnant 
and weighed 350g. The nursing nest 
was found on 27 July and seven young 
(6 males, 1 female) were marked. Off-
spring was blind and had only a downy 
fur cover emerging, typical for 10-14 
day-old squirrels (Eibl-Eibe-sfeldt, 
1951; Tittensor, 1977). A litter of seven 
was the largest one ever recorded for 
red squirrels (but see Ognev, 1940). In 
the UK, average litter size between 
birth and first weeks of life was 3.2 
young (N = 81, Tittensor, 1977), and 
reviews reported litter sizes ranging 
from 2-5 (Lange et al., 1994) and 1-4, 
max. 6 young (Tittensor, 1977; Lurz et 
al., 2005). We only found a single cita-
tion from the (translated) Russian lit-
erature on “10 squirrels found in a 
nest” (in Ognev, 1940), but it is not 
clear whether this refers to a single lit-
ter. Detailed capture-mark-recapture 
studies in Western Europe reported 
maximum litter size (between birth and 
1-week old) of four and average litter 
size of 3.0 ± 0.9 (N = 9, Wauters and 
Lens, 1995). Finally, even in a pine 
wood with supplementary feeding, av-
erage litter size was not higher than in 
natural populations (mean ± S.D. = 
3.17 ± 0.47 and 2.40 ± 0.31 young in 
63
Mari et al. 
 
two consecutive years, Shuttleworth, 
1996 cited in Lurz et al., 2005). Off-
spring losses during lactation can be 
high, and even complete litters can be 
lost due to predation or when body 
condition of the mother is reduced (loss 
of body mass between birth and wean-
ing, Wauters and Dhondt, 1989, 1995; 
Wauters and Lens, 1995; Lurz et al., 
2005). In the case of the litter of seven, 
two juveniles were observed at less 
than 50m from the nest tree on August 
31st and September 1st (ca. 6-7 weeks 
old), foraging on pine cones. One of 
them, a male, was recaptured at the end 
of October (ca. 3.5 months old), when 
also a male of the litter of three was 
trapped (nearly 6 months old). 
Finally, trapping revealed that the other 
two resident adult females (F2359, 
body mass 295g, and F2429, body 
mass 395g) also produced a litter 
(caught in lactation on August 31st and 
September 2nd , respectively) and three 
juveniles were caught and marked on 1 
and 2 September 2007 when about 10-
12 weeks old (200 – 230g). One of the 
females probably had two litters in 
2007, since we also captured a small 
juvenile (100g, ca. 6-7 weeks old) on 
25 June 2007. This will be tested using 
parent assignment tests on DNA sam-
ples. Overall offspring sex-ratio (10 
males, 4 females) was not different 
from even (χ21 = 2.57; P = 0.11). Body 
mass differed strongly among breeding 
females (range 295 – 395g), but two of 
them had a high lactation mass for fe-
male squirrels in populations through-
out the Alps (Wauters et al., 2007). 
How can we explain this extremely 
high reproductive rate (14 offspring 
from 4 adult females = 3.5 
juv./reproducing female)? We suggest 
that in this extreme habitat, with strong 
annual fluctuations in seed-crop size 
and thus food availability and long and 
cold winters with deep snow-cover, 
winter mortality can be very high. This 
was confirmed by our preliminary data 
(winter survival September 2006 - 
March 2007 = 0.25). Hence, the prob-
ability that squirrels only have one or 
two years to reproduce is high and fe-
males should adopt a trade-off between 
high reproductive investments when 
food abundance is high, against re-
duced survival (Williams, 1966; Clut-
ton-Brock, 1988). We therefore predict 
that in mountain pine forests, red squir-
rels are selected to reproduce as year-
lings and to invest heavily in reproduc-
tion in the first year after birth they 
have access to a rich seed-crop. To test 
this hypothesis, we will continue to 
monitor annual seed-crop production, 
squirrel numbers, reproductive success 
of all resident females and offspring 
survival/recruitment over the following 
years. 
 
ACKNOWLEDGEMENTS 
 
We thank AEM company for logistic 
support, and Matteo Wauters for 
company and tree climbing. Construc-
tive criticism by two referees helped to 
improve the manuscript. This study is 
part of the ASPER (Alpine Squirrel 
Population Ecology Research) project, 
funded by the province of Sondrio 
(Servizio Agricoltura, Caccia e Pesca, 
Settore Risorse Ambientali) and the 
Stelvio National Park to Istituto Oikos 
NGO, Milan. Additional financial 
support was given by MIUR (Ministero 
dell’Istruzione, dell’Università della 
ricerca, project COFIN 2003, number 
64
Record litter size in red squirrel 
 
2003053710-006) to Insubria Universi-
ty of Varese. This is paper12 of the 
ASPER project. 
 
REFERENCES 
 
Boutin S., Wauters L.A., McAdam A.G., 
Humphries M.M., Tosi G. and Dhondt 
A.A. 2006. Anticipatory reproduction 
and population growth in seed preda-
tors. Science, 314: 1928-1930. 
Clutton-Brock T.H. 1988. Reproductive 
success, University of Chicago Press, 
Chicago. 
Eibl-Eibesfeldt I. 1951. Beobachtungen zur 
Fortplanzungsbiologie und Jugendent-
wicklung des Eichhörnchens (Sciurus 
vulgaris L.). Z. Tierpsychol., 8: 370-
400. 
Humphries M.M. and Boutin S. 2000. The 
determinants of optimal litter size in 
free-ranging red squirrels. Ecology, 81: 
2867-2877. 
Lange R., Twisk P., Van Winden A. and 
van Diepenbeek A. 1994. Zoogdieren 
van West-Europa, KNNV-Uitgeverij, 
Utrecht, pp. 88-92. (in Dutch) 
Lurz P.W.W., Gurnell J., and Magris L. 
2005. Sciurus vulgaris. Mammalian 
Species, American Society of Mamma-
logists, 769: 1-10 
Ognev S. I. 1940. Mammals of the 
U.S.S.R. and adjacent countries. Mam-
mals of eastern Europe and northern 
Asia. Translated from Russian, Israel 
programme for scientific translations, 
Jerusalem, 1966, Vol IV: 284-377.  
Reznick D. 1992. Measuring the cost of 
reproduction. Trends Ecol. Evol., 7: 
42-45. 
Tittensor A.M. 1977. Red Squirrel. In: 
Corbet G.B., Southern H.N (eds.), The 
Handbook of British Mammals, 2nd 
edition. Blackwell Scientific Publica-
tions, Oxford, 153-164. 
Wauters L.A. and Dhondt A.A. 1989. Body 
weight, longevity and reproductive 
success in red squirrels (Sciurus vulga-
ris). J. Anim. Ecol., 58: 637-651. 
Wauters L.A., and Dhondt A.A. 1993. Im-
migration patterns and success in red 
squirrels. Behav. Ecol. Sociobiol., 33: 
159-167. 
Wauters L.A. and Dhondt A.A. 1995. 
Components of lifetime reproductive 
success of female Eurasian red squir-
rels. Oikos, 72: 402-410. 
Wauters L.A. and Lens L. 1995. Effects of 
food availability and density on red 
squirrel (Sciurus vulgaris) reprodu-
ction. Ecology, 76: 2460-2469. 
Wauters L.A., Bertolino S., Adamo M., 
Van Dongen S. and Tosi G. 2005. 
Food shortage disrupts social organiza-
tion: the case of red squirrels in conifer 
forests. Evol. Ecol., 19: 375-404. 
Wauters L.A., Bijnens L. and Dhondt A.A. 
1993. Body mass at weaning and juve-
nile recruitment in the red squirrel. J. 
Anim. Ecol., 62: 280-286. 
Wauters L.A., Vermeulen M., van Dongen 
S., Bertolino S., Molinari A., Tosi G. 
and Matthysen E. 2007. Effects of spa-
tio-temporal variation in food supply 
on red squirrel Sciurus vulgaris body 
size and body mass and its conse-
quences for some fitness com-ponents. 
Ecography, 30: 51-65. 
Weiner J. 1992. Physiological limits to sus-
tainable energy budgets in birds and 
mammals: ecological implications. 
Trends Ecol. Evol., 7: 384-388. 
Williams G.C. 1966. Natural selection, the 
costs of reproduction, and a refinement 
of Lack’s principle. Am. Nat., 100: 
687-690. 
 
 
 
65


Record litter size in the Eurasian red squirrel (Sciurus vulgaris)

&lt;strong&gt;Abstract&lt;/strong&gt;&#13; In the literature, litter size in red squirrels &lt;em&gt;Sciurus vulgaris&lt;/em&gt; ranges between 1-6 offspring. Here we report a record litter size, in Europe, of 7 young (6 males, 1 female), marked in the nest when 10-14 days old, in a high-elevation mountain pine habitat, Lombardy Alps (Northern Italy). We discuss implications of extreme winter conditions for life-history strategies and possible trade-offs between high reproductive investment and short life-span.&#13; &#13; &lt;strong&gt;Riassunto&lt;/strong&gt;&#13; &lt;strong&gt;Dimensione eccezionale della cucciolata di scoiattolo comune (&lt;em&gt;Sciurus vulgaris&lt;/em&gt;).&lt;/strong&gt;&#13; Dai dati di letteratura, il numero di esemplari per cucciolata nello scoiattolo comune &lt;em&gt;Sciurus vulgaris&lt;/em&gt; varia generalmente da 1 a 6. Nel presente lavoro viene riportata per la prima volta in Europa la segnalazione di una cucciolata di 7 esemplari (6 maschi e 1 femmina), che sono stati marcati nel nido all’età di 10-14 giorni. Il nido è stato individuato in un habitat forestale dominato da pino mugo nel Parco Nazionale dello Stelvio, in Lombardia. Vengono prese in esame le implicazioni che le condizioni estreme, nel periodo invernale, possono avere sulle strategie riproduttive dello scoiattolo comune

Veronica Mari

Stefania Martini

Claudia Romeo

Ambrogio Molinari

Adriano Martinoli

Guido Tosi

Lucas Wauters

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Record litter size in the Eurasian Red Squirrel (&lt;em&gt;Sciurus vulgaris&lt;/em&gt;)

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Record litter size in the eurasian red squirrel (Sciurus vulgaris)

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